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Systematic Botany (2018), 43(2): pp. 485–496
© Copyright 2018 by the American Society of Plant Taxonomists
DOI 10.1600/036364418X697210
Date of publication June 21, 2018
Reinstatement of the Genus Piofontia: A Phylogenomic-based Study Reveals the Biphyletic
Nature of Diplostephium (Asteraceae: Astereae)
Oscar M. Vargas
Department of Ecology and Evolutionary Biology, University of Michigan, 830 N. University Avenue, Ann Arbor,
Michigan 48109, USA
Integrative Biology and Plant Resources Center, The University of Texas at Austin, Austin, Texas 78712, USA
(oscarvargash@gmail.com)
Communicating Editor: Chrissen E. C. Gemmill
Abstract—A recent phylogenomic study has shown that Diplostephium in its broad sense is biphyletic. While one of the clades comprises 60 species
distributed mainly in the Northern Andes, the clade that contains the generic type, Diplostephium ericoides,contains48species,andprimarilyinhabitsthe
Central Andes. Here, I propose to reinstate the generic name Piofontia and transfer to it the species of Diplostephium in the Northern Andean clade. Piofontia
consists, then, of 60 species of woody subshrubs, shrubs, and small trees inhabiting high Andean forests and p´aramos of Costa Rica, Colombia, Venezuela,
and Ecuador. A morphological description is provided for the genus Piofontia along with a species list with 60 new combinations. Dysaster cajamarcensis is
shown to be a synonym of Diplostephium serratifolium. Finally, a brief discussion about the morphological evolution of South American Astereae is provided.
Keywords—Andes, South America, p´aramo.
Kunth (1820) proposed Diplostephium Kunth with a single
species, D. lavandulifolium Kunth (5D. ericoides (Lam.) Cab-
rera), and defined the genus with the following diagnostic
characteristics: branched shrubs with dense foliage, alternate
linear leaves, solitary capitula, hemispherical involucres with
numerous imbricate phyllaries, epaleate foveolate receptacles,
radiate heterogamous capitula, tubular hermaphrodite disk
florets, peripheral ray florets, and double pappus with a short
exterior row of scale-like bristles and an inner row of longer
barbellate bristles. Weddell (1855) reinterpreted Diplostephium,
adding a geographic component to his concept by defining the
genus as Andean shrubs of montane habitats with alternate
and often tomentose leaves, terminal solitary capitula on
branchlets or in a corymb, foveolate receptacles, and white or
purple rays. Weddell’s (1855) definition expanded the mor-
phological boundaries of Diplostephium and demarcated its
geographical distribution. Weddell (1855) also described 11
new taxa, transferred five species into the genus, and proposed a
subgeneric division of two groups: plants with solitary capitula
and plants with capitula borne in a corymb. Hieronymus (1894,
1896, 1900, 1905) added ten taxa to the genus, five of which are
now considered synonyms of previously described species
(Vargas 2011).
In the twentieth century, Blake described 28 new taxa for
Diplostephium and published two major revisions. In his first
revision, Blake (1922) proposed a subgeneric classification of
five series based on foliar and floral characters, partially fol-
lowing the subgeneric division proposed by Weddell (1855). In
the second revision, Blake (1928) recognized a total of 43
Diplostephium species but reduced the number of series to
three. Subsequently, Cuatrecasas (1943b, 1969) added nu-
merous names to Diplostephium and published two compre-
hensive studies of the genus. In his second work, Cuatrecasas
(1969) dealt only with the Colombian species and proposed a
generic subdivision of 12 series for Diplostephium consisting of
Blake’s (1922) original five plus seven new series. Cuatrecasas
(1969) listed 53 species for Colombia and estimated a total of 90
species for the genus. Although some species have been added
to Diplostephium since 1969, Cuatrecasas’(1969) study is the
most comprehensive taxonomic work of the genus to date. In
its recent circumscription (Vargas and Madri~
n´an 2006; Vargas
2011), Diplostephium s. l. comprises 111 species distributed in
the mountains of Central and South America in Costa Rica,
Colombia, Venezuela, Ecuador, Peru, Bolivia, and northern
Chile.
A recent phylogenomic study (Vargas et al. 2017) showed that
Diplostephium sensu Cuatrecasas (1969) is biphyletic, based on
a double-digest restriction site-associated DNA sequencing
(ddRAD) tree obtained by the authors (Fig. 1). A monophyletic
group of Diplostephium s. l. species, distributed mainly in the
Northern Andes, is sister to a clade that comprises Blakiella
Cuatrec., Hinterhubera Sch.Bip. ex Wedd., and Laestadia Kunth ex
Less. A second group of mainly Central Andean species,
Diplostephium s. s., forms a clade with Parastrephia quadrangularis
(Meyen) Cabrera. The Central Andean clade is nested in a clade
with Archibaccharis Heering, Aztecaster G.L.Nesom, Baccharis L.,
Exostigma Sancho, Floscaldasia Cuatrec., Heterothalamus Less.,
Laennecia Cass., Lagenophora Cass., and Westoniella Cuatrec. When
the Diplostephium s. l. species not sampled by Vargas et al. (2017)
are morphologically assigned to the two recovered clades (see
below), the Northern Andean and the Central Andean clades
contain 60 and 48 species, respectively.
Because the nomenclatural type of Diplostephium,D. ericoides
(Lam.) Cabrera, is part of the Central Andean clade, the correct
genericnameforthespeciescomprisingtheNorthernAndean
clade is Piofontia Cuatrec. Cuatrecasas (1943a) proposed Piofontia
as a monotypic genus consisting of P. colombiana Cuatrec. but
later (Cuatrecasas 1953) transferred P. colombiana into Diplo-
stephium as D. colombianum (Cuatrec.) Cuatrec. The previous
Cuatrecasas’series, erected primarily for the Colombian species,
are not recognized here because, for the most part, they do not
correspond to natural groups in either Piofontia and Diplo-
stephium. In addition to the monotypic series, the only mono-
phyletic series are Diplostephium ser. Denticulata, albeit with low
support (clade 1, Fig. 1), and Diplostephium ser. Schultziana if
Piofontia apiculata is included in the series (clade 2, Fig. 1).
Finally, I propose Dysaster cajamarcensis H.Rob. & V.A.Funk
as a taxonomic synonym of Diplostephium serratifolium Cuatrec.
based on morphology and the phylogeny of Vargas et al. (2017).
Taxonomic Treatment
PIOFONTIA Cuatrec. Caldasia 2: 5. 1943. TYPE:Piofontia colombiana
Cuatrec., Caldasia 2: 5. 1943.
Small trees, shrubs or subshrubs 0.1–10 m tall, woody,
branching sympodial by substitution with branches
485
terminated by capitulescences. Branches cylindrical, minutely
ribbed, tomentose or glabrous, glandular or eglandular, striate
when old; terminal shoots often tomentose. Leaves alternate
with phyllotaxis of five, petiolate, pseudopetiolate, or sessile,
often mucronate; lamina 3–230 30.7–85.0 mm, linear, lan-
ceolate, ellipsoid, oblong, ovate, or obovate; margins entire,
denticulate, or serrate, membranous to coriaceous, usually
revolute, sometimes flat, often with adaxial and abaxial
Fig. 1. Cladogram of Diplostephium,Piofontia, and their allies obtained with nuclear double-digest restriction site-associated DNA sequencing (ddRAD)
by Vargas et al. (2017) using maximum likelihood. Edges with bootstrap support ,90 are dashed. Letters next to species indicate the subtribe to which the
genus belongs: B 5Baccharidinae, H 5Hinterhuberinae, L 5Lagenophorinae, and P 5Podocominae. 1) Indicates the Denticulata clade, and 2) indicates the
Schultziana clade (both defined in the main text). The section mark (§) indicates the type species of Diplostephium. The map shows the type localities for
the species of Diplostephium (diamonds) and Piofonta (triangles). Figure modified from Vargas et al. (2017).
SYSTEMATIC BOTANY [Volume 43486
surfaces of a different color; adaxial surface often lanate when
young and glabrous when old, glandular or eglandular, with
central vein impressed and canaliculate, secondary venation
usually conspicuous in leaves wider than 12 mm and tertiary
venation usually conspicuous in leaves wider than 15 mm;
abaxial surface often densely lanate, whitish, yellowish, or
ocherous, central vein prominent, with secondary and tertiary
venation impressed or inconspicuous.
Capitulescence of terminal solitary or multiple heads
arranged in corymbs, racemes, or umbels. Capitula heterog-
amous, radiate, rarely disciform; involucre 4–15 mm long,
3–15 mm diam, tubular, cupulate, campanulate, or subconical,
3–7 seriate; phyllaries numerous, imbricate, unequal,
0.5–12.0 30.4–2.8 mm, ovate to linear, semicoriaceous or
coriaceous, often dorsally lanate and colored towards the apex.
Ray florets 5–140, with corolla 2.5–15.0 mm long, white to
purple; tube 1–6 mm long, usually papillose-pilose, rarely with
one or two lobes opposite to the limb; limb 0.5–12.0 3
0.4–2.0 mm, linear, oblong-elliptic, oblong-ovate, oblong-
obovate, with 3–4 veins; lobes (2–)3(–4), 0.1–0.8 mm long,
triangular and unequal; stigmatic branches 0.2–2.0 mm long,
linear or subulate with papillose margins; ovary 0.6–5.0 mm
long, glabrous or pilose, oblong, ovate, obovate, or oblong-
ellipsoid, ribbed, glandular or eglandular, always ovulate and
fertile; pappus biseriate, straw-colored, reddish, or purplish,
outer bristles 0.1–5.0 mm long, filiform and barbellate, interior
bristles 2.0–8.0 mm long, barbellate, often with flattened and
widened apex. Disk florets 3–114, staminate, corollas actino-
morphic, 3.0–9.0 mm long, tubular, tubular-campanulate, or
tubular-infundibuliform, whitish, green, yellow, or purple,
usually papillose-pilose, 5-lobed; tube 2.0–6.0 mm long; throat
1.0–4.0 mm long, usually papillose-pilose; lobes triangular
0.4–2.6 mm long, usually with papillose apex; anthers
1.0–2.5 mm long, oblong, briefly auriculate at the base, apical
appendix membranous, triangular, and obtuse; stigmatic
branches 0.3–1.2 mm long, linear or lanceolate, exteriorly
papillose; ovary 0.8–7.0 mm long, linear or oblong, ribbed,
glabrous to pilose, glandular or eglandular; pappus biseriate,
straw-colored, reddish, or purplish, outer bristles always fi-
liform, usually barbellate, 0.1–4.5 mm long, interior bristles
3–8 mm long, usually barbellate, often with flattened and
widened apex. Receptacle 1–5 mm in diam, alveolate, often
muricate.
Piofontia currently consists of 60 species and is a major
component of the flora of the Northern Andes, the Sierra
Nevada of Santa Marta, and the Talamanca Cordillera. The
southern boundary of Piofontia is central Ecuador, near the
Huancabamba depression, which defines the southern bound-
ary of the Northern Andes (Weigend 2004; Luebert and Pliscoff
2006; Luebert and Weigend 2014). Most o f the species inhabit the
p´aramo ecosystem, but a clade of approximately 16 species,
which contains the species of Diplostephium ser. Denticulata sensu
Cuatrecasas (1969), occurs in the cloud forest (clade 1, Fig. 1).
I propose the following new combinations, taking into ac-
count the evidence discussed above. Morphological evidence
(Cuatrecasas 1969; Vargas and Madri~
n´an 2006) was employed
to place the species not sampled by Vargas et al. (2017) in
Piofontia or Diplostephium. Finally, I describe two clade names
within Piofontia to facilitate communication in future studies
in the genus.
1. Piofontia alveolata (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium alveolatum Cuatrec., Caldasia 2: 224. 1943.
TYPE:COLOMBIA. Boyac ´a: p ´aramo del Alto del Escobal,
entre Soat´a y Cocuy, 3800–3900 m, arbolito 2–4 m, Sep 15
1938, Cuatrecasas & Garc´
ıa-Barriga 1758 (holotype: COL!,
isotypes: BC, F!, P, US!).
2. Piofontia anactinota (Wedd.) O.M.Vargas, comb. nov.
Diplostephium anactinotum Wedd., Chlor. And. 1: 201.
1856. TYPE: COLOMBIA. Magdalena: Sierra Nevada de Santa
Marta, fr´utex, flores albi, 1843, Funck 390 (lectotype: P,
isotypes: G, P).
3. Piofontia antioquensis (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium antioquense Cuatrec., Proc. Biol. Soc. Wash.
74: 12. 1961. TYPE: COLOMBIA. Antioquia: Medell´
ın, monte
El Boquer´on, Alto de los Bald´
ıos, p´aramo 3150 m, bosque
andino fragmentario marginal, ´arbol 4–5 m, hoja cra-
si ´uscula, verde amarillenta oscura haz, blanquecina
env´es, inflorescencia blanquecina, borde de las br´acteas
involucrales algo pardusco, l´
ıgulas blancas, Apr 9 1958,
Cuatrecasas, Llano & Gutierrez 24226 (holotype: US!, iso-
types: B, F!, G, MEDEL, NY, P, US!).
4. Piofontia apiculata (S.F.Blake) O.M.Vargas, comb. nov.
Diplostephium apiculatum S.F.Blake, Proc. Biol. Soc. Wash.
49: 79. 1936. TYPE: COLOMBIA. Norte de Santander: P ´aramo
de Santurb´an, entre Tona y Mutiscu´a, 3900 m, matita
le~
nosa, l´
ıgulas blancas, fl ´osculas verdes, Jan 19 1927, Killip
& Smith 19571 (holotype: US!, isotypes: A, COL!, GH, NY).
5. Piofontia bicolor (S.F.Blake) O.M.Vargas, comb. nov.
Diplostephium bicolor S.F.Blake, Contr. U. S. Natl. Herb. 24:
85. 1922. TYPE: COLOMBIA. Cauca: Central Cordillera head
waters of R´
ıoPalo, Tierra Adentro, 2500–3000 m, small
tree 2–4 m, Jan 1906, Pittier 1084 (holotype: US!, isotypes:
GH, F!).
Diplostephium tabanense Cuatrec., Caldasia 2: 216. 1943. TYPE:
COLOMBIA. Nari~
no: P´aramo del T ´abano, alto de la cor-
dillera entre Pasto y El Encano, vert occid., 3200 m,
arbolito, Jan 11 1941, Cuatrecasas 11932 (holotype: COL!,
isotypes: BC, F!, US!).
6. Piofontia camargoana (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium camargoanum Cuatrec., Phytologia 23: 351.
1972. TYPE: COLOMBIA. Boyac´a: Arcabuco, alrededores de la
poblaci´on 2739–2850 m, Oct 20 1965, Huertas & Camargo
6309 (holotype: US!, isotype: COL!).
7. Piofontia cayambensis (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium cayambense Cuatrec., Brittonia 8: 183. 1956.
TYPE: ECUADOR. Napo-Pastaza: p´aramo NE Volc ´an Cayambe,
12,500 ft, woody Compositae on moist p´aramo and ravines
leading to alpine lake, Feb 10 1953, Prescott 329 (holotype:
NY).
8. Piofontia chrysotricha (S.D´
ıaz & B.L.Restrepo) O.M.Vargas,
comb. nov. Diplostephium chrysotrichum S.D´
ıaz & B.L.
Restrepo., Revista Acad. Colomb. Ci. Exact. 19: 243. 1994.
TYPE: COLOMBIA. Tolima: Cajamarca, corregimiento de
Anaime, P´aramo de los Valles, La Cascada, Hacia Santa
Helena, 3710 m, Arbolito 2.5 m alt, 3 cm di´am., l´
ıgulas
blancas, flores vino tinto, en turbera, hojas con olor a
mentol, pegajosas al tacto, Jun 1994, Restrepo 371 (holo-
type: COL!, isotype: US!).
9. Piofontia cinerascens (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium cinerascens Cuatrec., Caldasia 3: 422. 1945.
VARGAS: REINSTATEMENT OF PIOFONTIA 4872018]
TYPE: COLOMBIA. Valle del Cauca: Cordillera Occidental,
Los Farallones de Cali, filo de la cordillera, matorrales de
p´aramo en el cerro de La Torre, 4000 m, arbolito, hoja
verde claro, mate haz, blanquecina env´es, nervio medial
verdoso blanquecino, involucro viol´aceo sucio, l´
ıgulas
blancas, corolas centrales blanco verdosas, Oct 10 1944,
Cuatrecasas 17851 (holotype: VALLE, isotypes: COL!, F!, P,
US!).
9.1. Piofontia cinerascens subsp. puracensis (Cuatrec.) O.M.
Vargas, comb. nov. Diplostephium violaceum var. puracense
Cuatrec., Caldasia 3: 424. 1945. Diplostephium cinerascens
subsp. puracense (Cuatrec.) Cuatrec., Webbia 24: 138. 1969.
TYPE: COLOMBIA. Cauca: Cordillera Central, al sur del
Volc´an Purac´e, filo de la cordillera en San Francisco,
3400–3450 m, arbolito muy ramoso de hoja muy compacta
y redondeada, hoja verde vivo en la haz, blanco tomen-
toso en el env´es, involucro verdoso en la base, viol ´aceo en
el extremo superior, l´
ıgulas blancas, fl´osculos blanco-
verdosos, anteras viol´aceas, Aug 23 1943, Cuatrecasas
14602 (holotype: VALLE, isotypes: COL!, F!, P, US!).
Diplostephium cinerascens var. centrale Cuatrec., Caldasia 3: 423.
1945. TYPE: COLOMBIA. Cauca: Cordillera Central, cabe-
ceras del R´
ıo Palo, quebrada del R´
ıoL´opez, Alto del
Duende, matorrales y bosquecillo de p´aramo, 3300–3350 m,
arbolito, hoja verde gris´acea clara haz, blanquecina env ´es,
br´acteas involucrales viol´aceas, l´
ıgulas blancas, “guasg ¨u´
ın”,
Dic 1 1944, Cuatrecasas 18796 (lectotype: US!, isotypes:
A, COL!, F).
10. PIOFONTIA COLOMBIANA Cuatrec., Feddes Repert. Nov.
Regni Veg. 55: 153. 1953. Diplostephium colombianum
(Cuatrec.) Cuatrec., Caldasia 2: 5. 1943. TYPE: COLOMBIA.
Boyac´a: Sierra Nevada del Cocuy, alto valle de Las
Lagunillas, pendientes rocosas mas abajo de los arenales,
4300–4400 m, fr´utex achaparrado, l´
ıgulas blancas que al
secarse se vuelven rosado-amarillosas, Sep 12 1938, Cuatrecasas
&Garc
´
ıa-Barriga 1492 (holotype: COL!, isotypes: F!, US!).
11. Piofontia coriacea (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium coriaceum Cuatrec., Webbia 24: 123. 1969.
TYPE: COLOMBIA. Magdalena: Sierra Nevada de Santa
Marta, flanco occidental, p´aramo 3100 m, arbusto 3 m,
tomento amarillo-ferrug´
ıneo-p´alido, l´
ıgulas crema,
fl´osculos rojizos, Jan 30 1959, Romero-Casta~
neda 7162
(holotype: COL!, isotype: US!).
12. Piofontia costaricensis (S.F.Blake) O.M.Vargas, comb.
nov. Diplostephium costaricense S.F.Blake, Contr. U. S. Natl.
Herb. 24: 82. 1922. TYPE: COSTA RICA. San Jos´e: Cerro de la
Muerte, 3100 m, Jan 1987, Pittier 10459 (holotype: US!,
isotype: G).
13. Piofontia crassifolia (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium crassifolium Cuatrec, Bull. Torrey Bot. Club
80: 401. 1953. TYPE: COLOMBIA. Cesar: Sierra del Perij ´a,
10 km, 12 km al noreste de Manaure, 48 km al este de
Valledupar, 1 km de la frontera con Venezuela, p´aramo
3000 m, cap´
ıtulos blancos, Feb 5 1945, Grant 10865 (ho-
lotype: US!, isotypes: NY, VEN).
14. Piofontia cyparissias (Wedd.) O.M.Vargas, comb. nov.
Diplostephium cyparissias Wedd., Chlor. And. 1: 203.
1856. TYPE: COLOMBIA. Magdalena: Sierra Nevada de
Santa Marta, vertiente N, flores albi, [without date],
Funck 387 (holotype: P, isotypes: G, F! [fragment], US!
[fragment]).
15. Piofontia elliptica (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium ellipticum Cuatrec., Caldasia 2: 212. 1943.
TYPE: COLOMBIA. Norte de Santander: P ´aramo de Fontib ´on,
2600–2750 m, entre Pamplona y Chitag´a, arbolito, Oct 16
1941, Cuatrecasas, Schultes, &Smith 12336 (holotype: COL!,
isotype: F!, US!).
16. Piofontia eriophora (Wedd.) O.M.Vargas, comb. nov.
Diplostephium eriophorum Wedd., Chlor. And. 1: 206. 1856.
TYPE: COLOMBIA. Tolima: Monte Tolima, en el l´
ımite in-
ferior de la nieve, 1844, Goudot s.n. (holotype: P, isotypes:
F! [fragment], FI, G, GH, P).
17. Piofontia farallonensis (Cuatrec.) O.M.Vargas, comb.
nov. Diplostephium floribundum subsp. farallonense Cua-
trec., Caldasia 3: 423. 1945. Diplostephium farallonense
(Cuatrec.) Cuatrec., Webbia 24: 135. 1969. TYPE: COLOMBIA.
Valle del Cauca: Cordillera Occidental, Los Farallones de
Cali, filo de la cordillera, matorrales de p ´aramo en el cerro
de La Torre, 4000 m, arbolito, hoja cori ´acea, r´
ıgida, verde
claro haz, blanquecina env´es, involucro verdoso-
amarillento, l´
ıgulas blancas, fl ´osculos viol ´aceos, Oct 10
1944, Cuatrecasas 17855 (holotype: VALLE, isotype:
COL!, F).
18. Piofontia floribunda (Benth.) O.M.Vargas, comb. nov.
Linochilus floribundus Benth., Pl. Hartw. 203. 1845. Dip-
lostephium floribundum (Benth.) Wedd., Chlor. And. 1: 205.
1856. TYPE: COLOMBIA. Cauca: Popay´an, P´aramo de Gua-
nacas, [without date], Hartweg 1126 (holotype: NY, iso-
types: F!, K, LD, P).
Linochilus ochraceus Sch.Bip., as a synonym in Weddell Chl.
And. 1: 205. 1857. Nom. nud.
Diplostephium ochroleucum Klatt, Engl. Bot Jahrb. 8: 37. 1886.
Type: Colombia. Cauca: [illegible] von Paletar´a, 3000 m,
Feb 5 1884, Lehmann 3579 (holotype: K, isotype: US!).
Aster ochroleucus (Klatt) Kuntze Rev. Gen. Pl. 3: 131. 1898.
19. Piofontia fosbergii (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium fosbergii Cuatrec., Bull. Torrey Bot. Club 80:
403. 1953. TYPE: COLOMBIA. Meta: Cordillera Oriental, R´
ıo
Arroz arriba de la confluencia con la Quebrada del
Pedregal, m´argenes con matorrales, 3445 m, arbusto 3 m,
cap´
ıtulos pardo verdosos, Aug 29 1943, Fosberg 20912
(holotype: F!, isotype: US!).
20. Piofontia frontinensis (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium frontinense Cuatrec., Revista Acad. Colomb.
Ci. Exact. 18: 123. 1991. TYPE: COLOMBIA. Antioquia: Mpio.
Urrao, P´aramo de Frontino, Llano Grande and hill to
north, small areas of dense mossy forest intermixed with
open Espeletia p´aramo, 3320–3450 m, shrub 1.5 m, rays
white disk dull gray, foliage sticky-glandular, Mar 2 1989,
McDougal, Rold´an, & Betancur 4424 (holotype: US!).
21. Piofontia glutinosa (S.F.Blake) O.M.Vargas, comb. nov.
Diplostephium glutinosum S.F.Blake, Proc. Biol. Soc. Wash.
49: 78. 1936. TYPE: COLOMBIA. Santander: P´aramo de los
Colorados, arriba de La Baja, 3900–4100 m, frutex
30 cm, br ´acteas con manchas purp´ureas, l´
ıgulas blancas,
fl´osculos con tubo amarillo verdoso y l ´obulos rosados,
SYSTEMATIC BOTANY [Volume 43488
estilos rosados, Jan 27 1927, Killip & Smith 18440 (holo-
type: US!, isotypes: A, COL! GH, K, NY).
22. Piofontia grantii (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium grantii Cuatrec., Bull. Torrey Bot. Club 80:
403. 1953. TYPE: COLOMBIA. Cesar: Sierra del Perij´a, 10 km
este-noreste de Manaure, 46 km al este de Valledupar,
3 km de la frontera con Venezuela, 2550 m, arbusto 2.6 m,
hojas pardo amarillentas env´es, l´
ıgulas blancas, Feb 4
1947, Grant 10791 (holotype: F!, isotype: NY, US!).
23. Piofontia heterophylla (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium heterophyllum Cuatrec., Caldasia 2: 230. 1943.
TYPE: COLOMBIA. Cundinamarca: Macizo de Bogot´a,
P´aramo de Cruz Verde, 3400–3500 m, ´arbolito acha-
parrado, 1–1.5 m, l´
ıgulas blancas, Sep 15 1940, Cuatrecasas
10457 (holotype: COL!, isotypes: BC, F!, G, P, US!).
24. Piofontia huertasii (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium huertasii Cuatrec., Webbia 24: 126. 1969.
TYPE: COLOMBIA. Cundinamarca: municipio de F´omeque,
p´aramo de Chingaza, finca la Laja, 3000 m, orillas del
camino, arbusto 2 m, hojas verdes opacas haz, estilos de
los fl´osculos del disco morados, Jan 10–18–20 1965,
Huertas & Camargo 5951 (holotype: US!, isotype: COL!).
25. Piofontia inesiana (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium inesianum Cuatrec., Webbia 24: 192. 1969.
TYPE: COLOMBIA. Magdalena: Sierra Nevada de Santa
Marta, flanco occidental, 3140 m, arbusto 4 m, Jan 29 1959,
Romero-Casta~
neda 7127 (holotype: COL!, isotype: US!).
26. Piofontia jaramilloi (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium jaramilloi Cuatrec., Phytologia 31: 317. 1975.
TYPE: COLOMBIA. Boyac´a: Cerro Berl´
ın, between Arcabuco
and la Palma (borderline between Boyac ´a and Santander),
in degraded Andean Forest, 2900 m, frutex 1.5–3 m tall,
leaves coriaceus, thick, yellowish green dull above,
ochraceous below, inflorescences and involucres ochra-
ceous, ligules white, disc corollas brownish, Mar 28 1973,
Cuatrecasas, Garc´
ıa-Barriga & Jaramillo 28667 (holotype:
US!, isotype: COL!).
27. Piofontia jenesana (S.D´
ıaz & M.E.Morales) O.M.Vargas,
comb. nov. Diplostephium jenesanum S.D´
ıaz & M.E.
Morales, Revista Acad. Colomb. Ci. Exact. 26: 7. 2002.
TYPE: COLOMBIA. Boyac´a: municipio de Jenesano, 2500 m,
arbusto, hojas verde intenso, flores blancas, [without
date], Molano 14 (holotype: UPTC, isotype: COL!).
28. Piofontia juajibioyi (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium juajibioyi Cuatrec., Webbia 24: 140. 1969.
TYPE: COLOMBIA. Norte de Santander: entre Chitag´ayEl
Cerrito, P´aramo del Almorzadero, un poco abajo (al sur)
del punto m´as ´alto del p ´aramo, 3900 m, fr ´utex 1 m, hojas
verde haz, casi blancas env´es, involucro verde velloso,
l´
ıgulas blancas, estrechas, vilano violeta-rojizo, flores de
disco verdes con ´apices purp ´ureos, Dic 31 1959 –Jan 1
1960, Barclay & Juajibioy 10388 (holotype: US!, isotypes:
COL!, F!, MO, U).
29. Piofontia julianii (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium juliani Cuatrec., Webbia 24: 127. 1969. TYPE:
VENEZUELA.T´achira, debajo del P´aramo de Tam´a, cerca
de la frontera con Colombia, faldas con bosque enano
ysubp´aramo, 2750–2950 m, shrubby vining, leaves
coriaceous, dark green above, buff brown below with
reticulate nerves, bracts brownish green, Jun 20–23 1967,
Steyermark & Dunsterville 98616 (holotype: US!, isotype:
NY).
30. Piofontia lacunosa (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium lacunosum Cuatrec., Bull. Torrey Bot. Club
80: 406. 1953. TYPE: COLOMBIA. Boyac´a: Sierra Nevada del
Cocuy, alto valle de Las Lagunillas, 4000–4300 m, arbusto
2–3 m, Sep 12 1938, Cuatrecasas 1443 (holotype: F!, iso-
types: COL!, US!).
31. Piofontia leioclada (S.F.Blake) O.M.Vargas, comb. nov.
Diplostephium leiocladum S.F.Blake, Amer. J. Bot. 15: 62.
1928. TYPE: COLOMBIA. Caldas: Cordillera Occidental,
Cerro Tatam´a zona de matorral al borde del p´aramo,
3300–3500 m, arbusto, l´
ıgulas blancas, Oct 8–10 1922,
Pennell 10531 (holotype: US!, isotypes: GH, PH).
32. Piofontia micradenia (S.F.Blake) O.M.Vargas, comb.
nov. Diplostephium micradenium S.F.Blake, Amer. J. Bot.
15: 49. 1928. TYPE: COLOMBIA. Caldas: Cordillera Occi-
dental, Cerro Tatam´a, 3300–3500 m, matorrales en
subp´aramo, l´
ıgulas blancas, flores de disco pardo-
amarillas, Oct 8 1922, Pennell 10533 (holotype: US!,
isotype: GH).
33. Piofontia mutiscuana (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium mutiscuanum Cuatrec., Webbia 24: 117.
1969. TYPE: COLOMBIA. Norte de Santander: entre Mutiscua
y Pamplona, 3400 m, shrub 10–12 ft, rays white, anthers
deep purple, pappus light brown, Feb 23 1927, Killip &
Smith 19708 (holotype: US!, isotypes: A, GH).
34. Piofontia nevadensis (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium nevadense Cuatrec., Brittonia 8: 182. 1956.
TYPE: COLOMBIA. Magdalena: Sierra Nevada de Santa
Marta, hasta 15,500 pies de altitud (4724 m), entre pe~
nas,
hasta el nivel de las nieves perpetuas, flor amarilla, Jan 13
1924, Wollaston 6 (holotype: K).
35. Piofontia oblongifolia (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium oblongifolium Cuatrec., Caldasia 2: 220.
1943. TYPE: COLOMBIA. Norte de Santander: cerro al noreste
de Pamplona, vertiente oriental, p´aramo entre matorrales
de bosque andino, 2770 m, arbolito, l´
ıgulas blancas, Sep 26
1940, Cuatrecasas & Garc´
ıa-Barriga 10238 (holotype: COL!,
isotypes: BC, F!, US!).
36. Piofontia obtusa (S.F.Blake) O.M.Vargas, comb. nov.
Diplostephium obtusum S.F.Blake, Contr. U. S. Natl. Herb.
24: 84. 1922. TYPE: VENEZUELA. Trujillo: P´aramo del Jab ´on,
3000–3200 m, Oct 2 1910, Jahn 24a (holotype: US!).
37. Piofontia ocanensis (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium ocanense Cuatrec., Brittonia 8: 182. 1956.
TYPE: COLOMBIA. Norte de Santander: entre Oca~
na y
Pamplona, 1878–1879, Kalbreyer 1199 (holotype: K).
38. Piofontia ochracea (Kunth) O.M.Vargas, comb. nov. Aster
ochraceus Kunth, Nov. Gen. Sp. Pl. 4: 94. 1820. Diplo-
stephium ochraceum (Kunth) Nees, Gen. Sp. Aster. 201.
1832. Tetramolopium ochraceum (Kunth.) DC., Prodr. 5: 262.
1836. TYPE: COLOMBIA. [erroneously cited as Ecuador.
Quito: monts de Quito, likely collected near Bogot´a,
Colombia], [without date], Humboldt & Bonpland s.n.
(holotype: P, F! [fragment]).
VARGAS: REINSTATEMENT OF PIOFONTIA 4892018]
Diplostephium denticulatum S.F.Blake, Contr. Gray Herb. 53: 25.
1918. TYPE: COLOMBIA. Cundinamarca: Macizo de Bogot´a:
Cerro de Guadalupe, 3000 m, Jul 1911, Apollinaire & Ar-
thur 11 (holotype: GH, isotype: US!).
39. Piofontia parvifolia (S.F.Blake) O.M.Vargas, comb. nov.
Diplostephium microphyllum Wedd., Chl. And. 1: 201. 1857.
Diplostephium parvifolium S.F.Blake, Contr. U. S. Natl.
Herb. 24: 74. 1922. TYPE: COLOMBIA [erroneously cited as
Venezuela]. Magdalena: Sierra Nevada de Santa Marta,
3000 m, arbustito, flores viol´aceas, [without date], Funck
388 (holotype: P, isotypes: GH [fragment], P [fragment]).
Linochilus microphyllus Sch.Bip., as a synonym in Wedd. Chl.
And. 1: 201. 1857. Nom. nud. Not Diplostephium micro-
phyllum (Vent.) Nees. Nom. illeg.
40. Piofontia perijaensis (S.D´
ıaz & G.P.M´endez) O.M.Vargas,
comb. nov. Diplostephium perijaense S.D´
ıaz & G.P.M´endez,
Revista Acad. Colomb. Ci. Exact. 21: 406. 1997. TYPE:
COLOMBIA. Cesar, Manaure, Serran´
ıa del Perij´a, camino
entre casa de vidrio y Cerro del Avi´on, 2900 m, 72°53’W
10°N, arbolito de 5 m, hojas ser´
ıceo-ferrug´
ıneas por el
env´es, verde n´
ıtido por la haz, br´acteas verdes variegadas
con morado, l´
ıgulas blancas, fl ´osculos pajizos, Nov 6 1993,
Rangel, Franco, Rudas, Olmos, Pardo y Clavijo 11212 (ho-
lotype: COL!, isotype: COL!).
41. Piofontia phylicoidea (Kunth) O.M.Vargas, comb. nov.
Aster phylicoides Kunth, Nov. Gen. Sp. Pl. 4: 93. 1820.
Tetramolopium phylicoides (Kunth) DC., Prodr. 5: 262. 1836.
Diplostephium phylicoides (Kunth) Wedd., Chlor. And. 1:
205. 1856. TYPE: COLOMBIA. [errouneously cited as Mexico,
see note below], [without date], Humboldt & Bonpland s.n.
(holotype: P “Herb. Bonpland”, isotype: P “Herb.
Bonpland”, F! [fragment]).
Linochilus phylicoides Sch.Bip., as a synonym in Wedd. Chl.
And. 1: 205. 1857. Nom. nud.
Aster crassifolius Klatt, Abhandl. Natur. Gesell. Halle 15: 326.
1882. TYPE: COLOMBIA. [without specific locality and date],
Linden 1 (lectotype: MA, isotypes: BM, F! [fragment],
LIL).
Diplostephium umbelliferum S.F.Blake, Contr. U.S. Nat. Herb.
24: 80. pl. 26. 1922. TYPE: COLOMBIA. Cundinamarca: Cerro
de Guadalupe, 1917, Ariste-Joseph s.n. (holotype: US!).
42. Piofontia pittieri (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium pittieri Cuatrec., Caldasia 2: 221. 1943. TYPE:
COLOMBIA. Valle del Cauca: Cordillera Central en el
macizo del Huila, P´aramo de Buena Vista, 3000–3600 m,
Jan 1906, Pittier 1174 (holotype: US!, isotype: F!).
43. Piofontia rangelii (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium rangelii Cuatrec., Phytologia 49: 74. 1981.
TYPE: COLOMBIA. Magdalena: Sierra Nevada de Santa
Marta, transecto de Buritaca, Filo la Cumbre, 3850 m,
arbustillo 1 m, l´
ıgulas blancas con tintes viol´aceos. Hojas
blancas por el env´es, Aug 19 1977, Rangel & Cleef 994
(holotype: COL!, isotypes: COL!, U, US!).
44. Piofontia revoluta (S.F.Blake) O.M.Vargas, comb. nov.
Diplostephium revolutum S.F.Blake, Contr. U. S. Natl. Herb.
24: 78. 1922. TYPE: COLOMBIA. Cundinamarca: Bogot ´a, 1917,
Ariste-Joseph A233 (holotype: US!).
Diplostephium revolutum var. longifolium Cuatrec., Trab. Mus.
Nac. Ci. Nat., Ser. Bot. 33: 134. 1936. TYPE: COLOMBIA.
Tolima: Vertiente sur del Nevado del Tolima, Las
Mesetas, 3800 m, “romero”, Jun 13 1932, Cuatrecasas 2892
(holotype: MA).
Diplostephium revolutum var. rubrum Cuatrec., Caldasia 2: 236.
1943. TYPE: COLOMBIA. Boyac´a: P´aramo de la Rusia, ver-
tiente SE, Boca de Monte, 3300–3400 m, arbusto, hojas
pegajosas, copa densa verde clara, Aug 4 1940, Cuatrecasas
10416 (holotype: COL!, isotypes: BC, F!, U, US!).
Diplostephium revolutum f. macrocephalum Cuatrec., Caldasia 2:
236. 1943. TYPE: COLOMBIA.Santander:P´aramo del Almor-
zadero vertiente norte, 3600–3800 m, arbolito, l´
ıgulas
blancas, Nov 28 1941, Cuatrecasas 13513 (holotype: COL!,
isotypes: BC, F!, US!).
45. Piofontia rhododendroides (Hieron.) O.M.Vargas, comb.
nov. Diplostephium rhododendroides Hieron., Bot. Jahrb.
Syst. 21: 340. 1896. TYPE: COLOMBIA. Nari~
no: Azufral de
T´uquerres, Laguna Verde, Jan 1840, Stubel 429 (holotype:
US! [fragment from destroyed specimen in B], F! [photo]).
Ibid: Volc´an Azufral, 3400–3500 m, escamas rojo carm´
ın
brillante, Feb 10 1972, Mora 1878A (topotype: US!).
Diplostephium cochense Hieron., Bot. Jahrb. Syst. 21: 341. 1896.
Type: Colombia. Nari~
no: llanura del R´
ıo Cocha con el
frailej´on, Aug 1869, Stubel 353 (holotype: US! [fragment
from destroyed specimen in B]).
46. Piofontia rhomboidalis (Cuatrec.) O.M.Vargas, comb.
nov. Diplostephium rhomboidale Cuatrec., Webbia 24: 151.
1969. TYPE: COLOMBIA. Boyac´a, Sierra Nevada del Cocuy,
alto valle de Las Lagunillas, 4000–4300 m, arbusto 2–3m,
l´
ıgulas azafrandas, vilanos rojizos, Sep 12 1938, Cua-
trecasas & Garc´
ıa-Barriga 1450 (holotype: US!, isotypes:
COL!, F!, P).
47. Piofontia ritterbushii (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium ritterbushii Cuatrec., Phytologia 23: 350.
1972. TYPE: COLOMBIA. Huila: west slope below Pico Norte
of Nevado del Huila, 4200 m (range is 4150–4250 m), very
abundant, Jan 11 1970, Ritterbush s.n. (holotype: US!).
48. Piofontia romeroi (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium romeroi Cuatrec., Webbia 24: 173. 1969. TYPE:
COLOMBIA. Magdalena: Sierra Nevada de Santa Marta,
flanco occidental, 3180 m, arbolito 5 m, flores rojas, Jan 29
1959, Romero-Casta~
neda 7115 (holotype: COL!, isotypes:
MO, US!, VEN).
49. Piofontia rosmarinifolia (Benth.) O.M.Vargas, comb. nov.
Linochilus rosmarinifolius Benth., Pl. Hartw. 197. 1845.
Diplostephium rosmarinifolium (Benth.) Wedd., Chlor. And.
1: 202. 1856. TYPE: COLOMBIA. Cundinamarca: Bogot´a,
alrederores, [without date], Hartweg 1092 (holotype: K,
isotypes: F!, G, LD, NY, P, US!, W).
Diplostephium baccharideum Blake, Contr. U. S. Natl. Herb. 24:
77. 1922. TYPE: COLOMBIA. Cundinamarca: Bogot´a, Mon-
serrate, 1917, Aristhe-Joseph B34 (holotype: US!).
Diplostephium rosmarinifolium var. baccharideum (Blake) Cua-
trec., Trab. Mus. Nac. Ci. Nat., Ser. Bot. 33: 135. 1936.
50. Piofontia rupestris (Kunth) O.M.Vargas, comb. nov. Aster
rupestris Kunth, Nov. Gen. Sp. Pl. 4: 94. Table 334. 1820.
SYSTEMATIC BOTANY [Volume 43490
Tetramolopium rupestre (Kunth) Ness, Gen. Sp. Ast. 203.
1832. Diplostephium rupestre (Kunth) Wedd., Chlor. And.
1: 206. 1856. TYPE: ECUADOR. Rucu Pichincha, [without
date], Humboldt & Bonpland 3047 (holotype: P “Herb.
Bonpl”, isotype: P “Herb. Bonpl”).
Aster pichinchensis Willd. ex Nees, Gen. Sp. Ast. 203. 1832.
Nom. nud.
Diplostephium mutisii Cuatrec., Trab. Mus. Nac. Ci. Nat., Ser.
Bot. 29: 25. 1935. TYPE: COLOMBIA. Tolima: p´aramo del
Nevado del Tolima, vert. S y SE, 4000–4300 m, Jun 15 1932,
Cuatrecasas 2882 (holotype: MA).
51. Piofontia santamartae (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium santamartae Cuatrec., Phytologia 52: 174. 1982.
TYPE: COLOMBIA. Magdalena: Sierra Nevada de Santa Marta,
“Transecto del R´
ıoBuritaca”, 3000 m alt, arbolito 3 m,
br´acteas e involucro de color vino tinto, hojas haz verdosas
env´es amarillo p´alido, Aug 1977, Rangel & Cleef 928 (ho-
lotype: COL!, isotypes: COL!, US! [fragment]).
52. Piofontia saxatilis (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium saxatile Cuatrec., Proc. Biol. Soc. Wash. 74:
14. 1961. TYPE: COLOMBIA. Magdalena: Sierra Nevada de
Santa Marta, flanco suroriental, hoya del R´
ıo Donach´
ı
entre la Laguna Esacuriba y unos enormes cantos y
pe~
nascos, 7870 m, ´arbolito 2–6 m, hoja blanda haz verde
clara, env´es ceniciento, involucro viol´aceo, corolas
tubulosas amarillentas, Oct 6 1959, Cuatrecasas & Romero-
Casta~
neda 24620 (holotype: US!, isotypes: COL!, F!, G, NY,
P).
53. Piofontia schultzii (Wedd.) O.M.Vargas, comb. nov.
Diplostephium schultzii Wedd., Chlor. And. 1: 204. 1856.
TYPE: COLOMBIA. Tolima: Volc´an Tolima, 9200 toeses, 21 fl.
Pourpres, Jan 1843, Linden 901 (holotype: P, isotypes: BM,
F!, G, K, P, W, US! [fragment]).
Linochilus jodopappus Sch.Bip. ex Wedd., Chl. And. 1: 204. 1857.
Nom. nud.
54. Piofontia tachirensis (V.M.Badillo) O.M.Vargas, comb.
nov. Diplostephium tachirense V.M.Badillo, Bol. Soc. Venez.
Ci. Nat. 10: 305. 1946. TYPE: VENEZUELA.T´achira, P´aramo
de Tam´a, Jul 15 1944, Steyermark 57360 (holotype: VEN,
isotypes: F!, NY, US!).
55. Piofontia tamana (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium tamanum Cuatrec., Caldasia 2: 214. 1943.
TYPE: COLOMBIA. Norte de Santander: P´aramo de Tam´a,
alrededores de La Cueva, residuos del bosque andino
l´
ımite, 3000–3200 m, l´
ıgulas blancas, fl´osculos verdosos,
Oct 28 1941, Cuatrecasas, Schultes, & Smith 12710 (holo-
type: COL!, isotypes: F!, P, S, U).
56. Piofontia tenuifolia (Cuatrec.) O.M.Vargas, comb.
nov. Diplostephium tenuifolium Cuatrec., Caldasia 2: 211.
1943. TYPE: COLOMBIA.Boyac´a, Valle de la Uvita, El
Hatico (entre Soata y Cocuy), bosque andino y mator-
rales, 3350 m, ´arbol grande, Sep 15 1938, Cuatrecasas &
Garc´
ıa-Barriga 1793 (holotype: COL!, isotypes: F!, P,
US!).
57. Piofontia tergocana (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium tergocanum Cuatrec., Webbia 24: 175.
1969. TYPE: COLOMBIA.Magdalena:SierraNevadade
Santa Marta, en las cabeceras del R´
ıo Sevilla, entre
grandes piedras en la base la subida al segundo pico al
este de la cabecera de la hoya, flanco occidental del
pico, 3770 m, “station 19”,arbustode3m,olorde
Espeletia, tallos inferiores desnudos, hojas verdes
haz, blanco lanosas env´es , br ´acteas involucrales
purp ´ureas, vilanos de blanco a rosado, muy resinoso,
pegajoso, Jan 28 1959, Barclay & Juajibioy 6762 (holo-
type: US!).
58. Piofontia venezuelensis (Cuatrec.) O.M.Vargas, comb.
nov. Diplostephium venezuelense Cuatrec., Caldasia 2: 233.
1943. TYPE: VENEZUELA.M´erida, Tabay, 2500–3000 m,
arbusto de 2–5 m de altura, hojas arom´aticas, cabezuelas
amarillo ocre, Nov 18 1930, Gehriger 473 (holotype: VEN,
isotypes: G, MO, NY, F!, US!).
59. Piofontia violacea (Cuatrec.) O.M.Vargas, comb. nov.
Diplostephium violaceum Cuatrec., Caldasia 2: 232. 1943.
TYPE: COLOMBIA. Caldas: Cordillera Central, vertiente oc-
cidental, vert. SW del Nevado del Ruiz, Termales, 3400 m,
arbusto de 2 m, o ´arbol hasta 8 m, flores centrales vio-
l´aceas, l´
ıgulas blancas, “romerillo,”Jun 4 1940, Cuatrecasas
9229 (holotype: COL!, isotypes: P, F!, US!).
60. Piofontia weddellii (S.F.Blake) O.M.Vargas, comb. nov.
Diplostephium weddellii S.F.Blake, Contr. U. S. Natl. Herb.
24: 79. 1922. TYPE: COLOMBIA. Guajira: Sierra Nevada de
Santa Marta, prov. del Riohacha, 4000–4400 m, arbusto,
flores amarillas, Mar 1 1852, Schlim 806 (holotype: P,
isotypes: BM, F!, G, K).
Diplostephium sessiliflorum Wedd., Chlor. And. 1: 204. Nom.
illeg. 1856. Not D. sessiliflorum Spreng. 1826.
Denticulata new clade name. Node based definition: the clade
originating with the most common ancestor of Piofontia
antioquensis, P. camargoana, P. huertasii, P. jaramilloi, P.
jenesana, P. mutiscuana, P. oblongifolia, P. ochracea, P.
tachirensis, and P. tenuifolia (clade 1, in Fig, 1). Name
originally proposed by Blake (1922) and later redefined by
Cuatrecasas (1969).
Schultziana new clade name. Node based definition: the clade
originating with the most common ancestor of Piofontia
alveolata,P. apiculata, P. costaricensis, P. juajibioyi, P.
rhomboidalis, and P. schultzii (clade 2, Fig. 1). Name orig-
inally proposed by Cuatrecasas (1969).
Additional Taxa—The following taxa described by earlier
studies as species of Diplostephium also belong in Piofontia, but
combinations have not been made here because they will
become taxonomic synonyms of recognized species of Piofontia
in the systematic treatment of the genus for Colombia (Vargas
in prep.)
1. Diplostephium dentatum Cuatrec.
2. Diplostephium fernandez-alonsoi S.D´
ıaz.
3. Diplostephium floribundum subsp. aequatoriense Cuatrec.
4. Diplostephium floribundum subsp. llanganatense Cuatrec.
5. Diplostephium floribundum subsp. putumayense Cuatrec.
6. Diplostephium floribundum subsp. cundinamarcense Cuatrec.
7. Diplostephium tolimense Cuatrec.
A New Synonym for Diplostephium serratifolium—In 2014,
Robinson and Funk (2014) erected the genus Dysaster H.Rob. &
V.A.Funk for a plant specimen with affinities to Diplostephium
(Robinson and Funk 2014: 35) but which they thought differed
VARGAS: REINSTATEMENT OF PIOFONTIA 4912018]
significantly from any species previously described in that
genus. Apparently, unknown to Funk and Robinson, a con-
specific from the same location (near Contumaz´a, Peru),
Diplostephium serratifolium Cuatrec., had been described by
Cuatrecasas (1982). Robinson and Funk (2014) focused their
diagnosis on the morphological features that, in their opinion,
supported the description of a new species and a genus: 1)
compressed achenes with only two ribs; 2) fully bisexual disk
florets with stigmatic lines; 3) involucral bracts narrowly
lanceolate, striped and pointy; 4) outer row of pappus dif-
ferentiated in short squamae; and 5) exserted inflorescence
with few heads borne by long peduncles. While all the
aforementioned characteristics match the description of D.
serratifolium made by Cuatrecasas (1982), these features are not
unique to Diplostephium serratifolium (5Dysaster cajamarcensis)
and therefore do not support the segregation of a new genus: 1)
the compression of the achenes is commonly not reported in
the description of new species and is mostly unknown in other
species of Diplostephium s. s., therefore it is premature to assign
this characteristic as diagnostic; 2) fertile ovaries in disk
flowers and stigmatic lines are also present in at least in D.
haenkei (DC.) Wedd. (Cuatrecasas 1982), and the state for this
character for numerous species is unknown; 3) striped narrow
involucre bracts are not uncommon in Diplostephium s. s. (e.g.
D. cajamarquillensis Cabr., D. callaensis Cuatrec.); 4) squamate-
like outer pappus is at least also present in D. ericoides and D.
azureum Cuatrec. and probably in other species too; and 5) long
peduncles are also present in D. wurdackii Cuatrec. Because of
the evidence explained above and the position of Diplostephium
serratifolium falling within the phylogenetic circumscription of
Diplostephium s. s. (Fig. 1), Dysaster cajamarcensis should be
considered a synonym of Diplostephium serratifolium.
DIPLOSTEPHIUM SERRATIFOLIUM Cuatrec. Phytologia 52: 176. 1982.
TYPE: PERU. Cajamarca: Contumaz´a, circa Contumaz´a,
2700 m, Jun 1960, Alza s.n. (holotype: LP, isotype: USM!,
US! [fragment, photo]).
Dysaster cajamarcensis H.Rob. & V.A.Funk. TYPE: PERU. Cajamarca:
Prov. Contumaz´a. 14 km S of Contumaz ´a on gravel road,
rocky slopes, western cordillera, evergreen forest, 2620 m,
Jul 1992, Stuessy, Crawford & Sagastegui 12686 (holotype:
US).
Artificial Key for the Identification of
Diplostephium
and
Piofontia
1. Ray corollas comparatively short, rarely long, 5–9(15) mm long. Subshrubs, shrubs, and small trees, 0.1–10.0 m tall, always with
determinate branches leading to a candelabrum-like branching pattern in which every branch is terminated by a capitulescence.
Leaves 0.3–23 cm long. Adaxial leaf surface usually eglandular or sparse-glandulose. Capitulescences of solitary capitula or with
up to 100 heads. Distribution: Costa Rica, Colombia, Venezuela, northern and central Ecuador (Azuay, Bolivar, Carchi,
Chimborazo, Cotopaxi, Imbabura, Napo, Pastaza, Pichincha, Tungurahua, Sucumbios) . . . .........................Piofontia
1. Ray corollas long, 8–22 mm long. Subshrubs and shrubs 0.2–3.0 m tall, with long indeterminate branches that bear short
branchlets topped with solitary capitula or a candelabrum-like branching pattern in which every branch is terminated by a
capitulescence. Leaves 0.2–8.0 cm long. Adaxial leaf surface usually eglandular or sometimes densely-glandulose. Capitulescences
comprised of solitary capitula or with up to 20 heads. Distribution: southern Colombia (Cauca, Huila, Nari~
no, Putumayo),
Ecuador, Peru, Bolivia, northern Chile (Arica y Parinacota, Tarapac´a, Antofagasta) . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplostephium
Discussion
Despite the overlapping morphological characteristics
among species of Piofontia and Diplostephium (microphyllous
leaves, heterogamous capitula, ray florets with a 2–3-lobed
limb, rays white to purple, and double pappus) a combination
of morphological characters can separate these clades
(Table 1). Piofontia species are characterized by being sub-
shrubs (0.10–0.49 m tall) or shrubs (0.5–3.9 m tall) with leaves
0.3–4.0 cm long, to small trees (4–10 m tall) with leaves 4–23 cm
long (Fig. 2). Subshrubs and small shrubs in Piofontia have
solitary or few (,20) capitula per capitulescence (Fig. 2E–G),
while taller shrubs and small trees always bear 20–100 heads
per capitulescence (Fig. 2D). The branching pattern of Piofontia
is very consistent, with flowers occurring on terminal branches
determining their growth (Fig. 2E); typically, three or four
branches develop from the axillary buds close to the capit-
ulescence, continuing the vertical growth of the plant. This
architecture gives Piofontia species a characteristic candela-
brum branching pattern (Fig. 2A–B). Ray corollas in Piofontia
are short or medium in length (Fig. 2D–G), 5–9 mm long,
Table 1. Comparative aspects between Diplostephium and Piofontia.
Piofontia Diplostephium
Habit Subshrubs, shrubs, or small trees up to 10 m tall. Subshrubs, shrubs up to 3 m tall.
Branching pattern Candelabrum-like, every branch is terminated by a capitulescence. Candelabrum-like, or long indeterminate branches
bearing short branches topped by solitary capitula.
Capitulescence Subshrubs bearing mostly solitary capitula. Shrubs and trees with
multiple heads per capitulescence. Number of capitula increases
with plant size and leaf width, up to 100 heads per capitulescence.
Subshrubs bearing mostly solitary capitula.
Shrubs bearing up to 20 heads per capitulescence.
Ray corollas length 5–9(15) mm 8–22 mm
Distribution Northern Andes, Sierra Nevada de Santa Marta, and Talamanca
Cordillera; Costa Rica, Colombia, Venezuela, northern and central
Ecuador (Azuay, Bolivar, Carchi, Chimborazo, Cotopaxi,
Imbabura, Napo, Pastaza, Pichincha, Tungurahua, Sucumbios).
Central Andes and southern Northern Andes;
southern Colombia (Cauca, Huila, Nari~
no,
Putumayo), Ecuador, Peru, Bolivia, northern Chile
(Arica y Parinacota, Tarapac´a, Antofagasta).
Habitat P´aramo and high Andean forest. Dry puna, humid puna, p´aramo, and upper boundary
of the high Andean forest.
Number of species 60 48
SYSTEMATIC BOTANY [Volume 43492
relative to Diplostephium with only some species reaching
lengths of 9–15 mm.
Diplostephium species are subshrubs or shrubs up to 3 m tall
(Fig. 3); subshrubs and medium-sized shrubs have leaves
0.2–3.0 cm long; scandent and large-sized shrubs have leaves
4–8 cm long. Most species have solitary capitula (Fig. 3A,
C–D), but scandent and large-sized shrubs exhibit up to 20
heads per capitulescence (Fig. 3B, E, F). A distinctive feature of
most Diplostephium species with solitary capitula is an archi-
tectural pattern in which long, indeterminate branches bear
short branchlets topped with solitary capitula (Fig. 3C–D). Ray
corollas in Diplostephium are always long, 8–22 mm in length
Fig. 2. A. Piofontia oblongifolia.B.P. eriophora.C.P. apiculata.D.P. camargoana.E.P. schultzii.F.P. rupestris.G.P. frontinensis. Notice the candelabrum-like
branching pattern in B and the short and medium length ray corollas in D–G.
VARGAS: REINSTATEMENT OF PIOFONTIA 4932018]
(Fig. 3D–G). Diplostephium now comprises 48 species that
inhabit the puna and the humid puna (high yunga) in Peru,
Bolivia, and northern Chile, and the p´aramo in Ecuador and
southern Colombia (where its distribution overlaps with
Piofontia). Diplostephium taxonomy remains largely unstudied
since the last revision of the genus that included Peruvian
species (Blake 1928); a morphological study is necessary to
properly define the morphological boundaries of the genus.
The ddRAD phylogeny of Vargas et al. (2017) places Par-
astrephia quadrangularis as sister to Diplostephium meyenii
Fig. 3. A. Diplostephium meyenii.B.D. haenkei.C.D. hartwegii.D.D. gnioides.E.D. barclayanum.F.D.lechleri.G.D. oxapampanum. Notice the short branches
topped with solitary capitula in C–D and the long ray corollas in D–G.
SYSTEMATIC BOTANY [Volume 43494
Wedd. nested within Diplostephium. Even though this po-
sition suggests that Parastrephia Nutt. species should be
transferred to Diplostephium, the phylogenetic position of
Parastrephia in the Vargas et al. (2017) phylogeny may be
biased by the high indices of hybridization and introgression
found among P. quadrangularis and D. cinereum,D. meyenii,
and D. sp. nov. CAJ2 (Vargas et al. 2017). Simulations have
shown that gene flow can affect phylogenetic topologies,
producing support for erroneous inferences of species trees
(Leach´e et al. 2014). Therefore, it is possible that Parastrephia
is an independent lineage from Diplostephium as suggested
by the nuclear ribosomal phylogeny of Vargas et al. (2017).
Parastrephia comprises three species of densely branched
cupressiform shrubs distinguished by having sessile mi-
crophyllous leaves, disciform heads, yellow corollas, pis-
tillate flowers in one series with tubular or limbate corollas,
disc flowers functionally staminate and tubular, and re-
ceptacles epaleate (Nesom 1993; Nesom and Robinson 2007).
Extended sampling that includes a broader Parastrephia
sampling is needed to resolve its position relative to
Diplostephium.
Incompatibility between the traditional classification of
Astereae (Nesom and Robinson 2007) and molecular phy-
logenies (Noyes and Rieseberg 1999; Sancho and Karaman-
Castro 2008; Brouillet et al. 2009; Karaman-Castro and
Urbatsch 2009; Sancho et al. 2010; Vargas and Madri~
n´an
2012; Vargas et al. 2017) demonstrates morphological la-
bility in the tribe and calls for a revaluation of its sub-
division. These results suggest that it is possible that similar
morphologies among genera in the tribe are the product of
parallel evolution, confounding morphological classifica-
tion relative to the true phylogeny of the tribe. In the specific
case of Diplostephium and Piofontia, the morphological
tendency to present a reduced vegetative morphology with
increasing altitude (shrubs/subshrubs, microphyllous
leaves), is likely the result of convergent adaptation to the
physiological stress produced by lower temperatures and
daily climate oscillations in the high Andes (Meinzer et al.
1994). Alternatively, these overlapping characters may be
of plesiomorphic nature. Future phylogenetic studies in
Astereae should focus on sequencing numerous nuclear loci
to obtain a robust phylogeny of the tribe as chloroplast and
mitochondrial phylogenies are often misleading in cases of
recent and ancient hybridization (Vargas et al. 2017). Recent
studies (Lagomarsino et al. 2014; Uribe-Convers and Tank
2016; Vargas et al. 2017) demonstrate the utility of
employing multiple loci and phylogenomic approaches in
systematics studies to elucidate novel taxa in biodiversity
hotspots.
Acknowledgments. I thank Jordan Bemmels, Kanchi Ganshi, Edg-
ardo Ortiz, Beryl Simpson, and Greg Stull for providing helpful comments
in early versions of this paper. I am grateful to the following herbaria for
providing resources and access to their collection: Museo de Historia
Natural de la Universidad de los Andes (ANDES), The Field Museum (F),
Herbario Universidad de Pamplona (HECASA), Herbario Sur Peruano
(HSP), Herbario Universidad de Antioquia (HUA), the University of
Michigan Herbarium (MICH), Herbario de la Pontificia Universidad
Cat´olica de Quito (QCA), the Plant Resources Center at The University of
Texas at Austin (TEX), the United States National Herbarium at the
Smithsonian Institution (US), Herbario Universidad Nacional Mayor de
San Marcos (USM). This study was supported by The University of Texas at
Austin (Plant Biology Program Awards, the C. L. Lundell Chair of Sys-
tematic Botany, The Linda Escobar Award), the Garden Club of America
(2012 Award in Tropical Botany), the Smithsonian Institution (Cuatrecasas
Award), and the National Science Foundation (postdoctoral support FESD
1338694 and DEB 1240869).
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